Diplonemidae

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Diplonemidae
Diplonema papillatum SEM image.gif
Scanning electron micrograph of Diplonema papillatum
Scientific classification
Domain:
(unranked):
Phylum:
Class:
Order:
Family:
Diplonemidae

Cavalier-Smith 1993 emend. Adl et al. 2019
Genus
Synonyms
  • Rhynchopodaceae Skuja 1948 ex Cavalier-Smith 1993

Diplonemidae is a family of biflagellated unicellular protists that may be among the more diverse and common groups of planktonic organisms in the ocean. Although this family is currently made up of three named genera; Diplonema , Rhynchopus , and Hemistasia , there likely exist thousands of still unnamed genera. [1] Organisms are generally colourless and oblong in shape, with two flagella emerging from a subapical pocket. [2] They possess a large mitochondrial genome composed of fragmented linear DNA. [3] These non-coding sequences must be massively trans-spliced, making it one of the most complicated post-transcriptional editing process known to eukaryotes.

Contents

Etymology

The word “Diplonemidae” come from the Greek words ‘diplo’, meaning two, and ‘nemat’, meaning thread. Together, Diplonemidae roughly translates to ‘two threads’, likely referring to the characteristic two flagella of the organism.

History of Knowledge

Primary studies done in the 1900s by Griessmann and Skuja had initially grouped Diplonemidae (or more specifically, the Diplonema and Rhynchopus taxa) with the euglenids. This was due to the two groups sharing many morphological similarities with the euglenids, such as metaboly, locomotion and a microtubule-reinforced feeding apparatus. [4] However, this conclusion was met with some controversy, as diplonemids lacked characteristic features shared by all euglenids, such as possession of pellicle strips and paraxonemal rods on their flagella. [5] Instead, they were placed in the euglenozoan phylum and shared this taxon with the kinetoplastids and euglenids. [6] The inclusion of Rhynchopus and Diplonema in the family Diplonemidae was later confirmed through nuclear 18S rRNA analysis. [7] As of now, Rhynchopus and Diplonema are part of the ‘classical’ diplonemid subgroup, and Hemistasia is part of the ‘nonclassical’ diplonemid subgroup.

Recently, there had been debates as to whether Diplonemidae were more closely related to the euglenids or kinetoplastids. It was not until analysis of cytosolic heat shock proteins that a sister relationship between the diplonemids and kinetoplastids was proposed. [5] This was further supported through phylogenetic analysis, which discovered that Diplonemidae possess a functional splice leader RNA that is characteristic of the kinetoplastids. [8] Additionally, the amino acid tryptophan is encoded by codon TGA in kinetoplastids and diplonemids, whereas in euglenids, the regular codon is used instead. [5]

Nevertheless, despite the similarities between kinetoplastids and diplonemids, the two taxa are still distinct form each other. Diplonemidae still lack a kinetoplast, and have a unique bi-flagellated trophic phase not seen in kinetoplastids. Presently, the kinetoplastids are regarded as the sister group to the Diplonemidae.

Although there are only three named genera within the Diplonemidae, environmental sequencing performed by the recent TARA Ocean Expedition concluded there are potentially thousands of genera, [1] suggesting that pelagic Diplonemidae are the most diverse planktonic eukaryotes in the oceans.

Habitat and Ecology

Although Diplonemidae are generally predators, some species display parasitic life strategies. [3] Diplonemidae display a rich diversity in marine and freshwater environments, with their relative abundance increasing with depth. [9] The diplonemids that exist in these different environments are genetically distinct, and exhibit slightly different lifestyles. The ‘classic’ diplonemids (i.e. Diplonema and Rhynchopus) are benthic, whereas the marine diplonemids, which include Hemistasia, are planktonic. [2]

There are potentially thousands of unknown marine Diplonemidae species, with this diversity highly stratified in accordance to depth. Although molecular sequencing confirms the existence of these unnamed marine Diplonemidae, information regarding their morphology and lifestyle is absent. [1] As marine Diplonemidae are the most abundant and genetically diverse protists (and potentially eukaryotes) in the sea, there are strong implications that they play a key role in aquatic ecosystems. As of now, this exact role is unknown.

Description of the Organism

Morphology and Anatomy

Classical diplonemids (i.e. Diplonema and Rhynchopus), are colourless and oblong in shape. They are approximately 20 μm in length and possess a microtubule layer underneath their plasma membrane. [10] Adjacent to it is a mitochondrion with discoidal cristae. [6] They also possess two flagella of equal length, both of which lack paraxial rods. [2] The two basal bodies originate from a subapical pocket, which merges with an adjacent feeding apparatus. [4] This feeding apparatus is surrounded by many food vacuoles and reinforced by microtubules. [10]

Nonclassical Diplonemidae (i.e. Hemistasia) are diverse in size but share many morphological aspects with the classic diplonemids. However, a great majority of these marine Diplonemidae have never been seen, with their existence only confirmed through molecular analysis. [2]

Although Diplonemidae do not possess pellicular strips like euglenids, they still move via metaboly. [3]

Diplonemidae also exhibit the compartmentalization of glycolytic and gluconeogenic enzymes into peroxisomes. These organelles are referred to as glycosomes, and is a characteristic feature also shared with their sister taxon, the kinetoplastids . [11]

Life cycles

Diplonemidae are capable of sexual reproduction, as genes involved in meiosis have been found. [12] Although marine diplonemids appear to reproduce sexually, not much is known about diplonemid reproduction as Euglenozoans rarely demonstrate sexual processes. [5]

Genetics

Diplonemidae have a very unique mitochondrial DNA arrangement. Although Diplonemidae possess a large mitochondrial genome, these do not contain any intact full-sized genes. Instead, their mitochondrial DNA consists of linear gene fragments of different sizes. Because each fragment is both full of repeats and incomplete, individually they are unable to code for a gene themselves. Instead, fragments are transcribed and spliced together using their own specialized trans splicing machinery. [3] Once spliced together, the transcript undergoes extensive editing to become recognizable RNA. This is accomplished by either Uracil-insertion, nucleotide deanimation, or substitution, which eventually generates a fully mature and translatable transcript. [13]

The Diplonemidae genome contains a spliced leader RNA gene, which confirms their use of mRNA spliceosome-dependent trans splicing during nuclear expression.

Diplonemids possess the gap3 gene which is specifically found in cyanobacteria and Pseudomonadota. This is likely due to lateral gene transfer following the divergence of diplonemids from the euglenoids. [14] As to date, this is one of the most-supported examples of lateral gene transfer from a bacterium to eukaryote and may have implications for diplonemid acquisition of biochemical abilities.

Taxonomy

Although presently made up of less than a dozen named species, the existence of thousands of unknown diplonemid species has been confirmed. [1]

Related Research Articles

<span class="mw-page-title-main">Euglenozoa</span> Phylum of protozoans

Euglenozoa are a large group of flagellate Discoba. They include a variety of common free-living species, as well as a few important parasites, some of which infect humans. Euglenozoa are represented by four major groups, i.e., Kinetoplastea, Diplonemea, Euglenida, and Symbiontida. Euglenozoa are unicellular, mostly around 15–40 μm (0.00059–0.00157 in) in size, although some euglenids get up to 500 μm (0.020 in) long.

<i>Euglena</i> Genus of unicellular flagellate eukaryotes

Euglena is a genus of single cell flagellate eukaryotes. It is the best known and most widely studied member of the class Euglenoidea, a diverse group containing some 54 genera and at least 200 species. Species of Euglena are found in fresh water and salt water. They are often abundant in quiet inland waters where they may bloom in numbers sufficient to color the surface of ponds and ditches green (E. viridis) or red (E. sanguinea).

<span class="mw-page-title-main">Euglenid</span> Class of protozoans

Euglenids are one of the best-known groups of flagellates, which are excavate eukaryotes of the phylum Euglenophyta and their cell structure is typical of that group. They are commonly found in freshwater, especially when it is rich in organic materials, with a few marine and endosymbiotic members. Many euglenids feed by phagocytosis, or strictly by diffusion. A monophyletic group consisting of the mixotrophic Rapaza viridis and the two groups Eutreptiales and Euglenales have chloroplasts and produce their own food through photosynthesis. This group is known to contain the carbohydrate paramylon.

<span class="mw-page-title-main">Kinetoplastida</span> Flagellated protists belonging to the phylum Euglenozoa

Kinetoplastida is a group of flagellated protists belonging to the phylum Euglenozoa, and characterised by the presence of a distinctive organelle called the kinetoplast, a granule containing a large mass of DNA. The group includes a number of parasites responsible for serious diseases in humans and other animals, as well as various forms found in soil and aquatic environments. The organisms are commonly referred to as "kinetoplastids" or "kinetoplasts".

<span class="mw-page-title-main">Excavata</span> Supergroup of unicellular organisms belonging to the domain Eukaryota

Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota. The group was first suggested by Simpson and Patterson in 1999 and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged. They are considered to be a basal flagellate lineage.

Trans-splicing is a special form of RNA processing where exons from two different primary RNA transcripts are joined end to end and ligated. It is usually found in eukaryotes and mediated by the spliceosome, although some bacteria and archaea also have "half-genes" for tRNAs.

<span class="mw-page-title-main">Discosea</span> Class of amoebae

Discosea is a class of Amoebozoa, consisting of naked amoebae with a flattened, discoid body shape. Members of the group do not produce tubular or subcylindrical pseudopodia, like amoebae of the class Tubulinea. When a discosean is in motion, a transparent layer called hyaloplasm forms at the leading edge of the cell. In some discoseans, short "subpseudopodia" may be extended from this hyaloplasm, but the granular contents of the cell do not flow into these, as in true pseudopodia. Discosean amoebae lack hard shells, but some, like Cochliopodium and Korotnevella secrete intricate organic scales which may cover the upper (dorsal) surface of the cell. No species have flagella or flagellated stages of life.

<span class="mw-page-title-main">Archamoebae</span> Phylum of protists

The Archamoebae are a group of protists originally thought to have evolved before the acquisition of mitochondria by eukaryotes. They include genera that are internal parasites or commensals of animals. A few species are human pathogens, causing diseases such as amoebic dysentery. The other genera of archamoebae live in freshwater habitats and are unusual among amoebae in possessing flagella. Most have a single nucleus and flagellum, but the giant amoeba Pelomyxa has many of each.

<i>Bodo saltans</i>

Bodo saltans is a free-living nonparasitic species of kinetoplastid flagellated phagotrophic protozoa that feed on bacteria. Bodo saltans cells have been reported in freshwater and marine environments.

<span class="mw-page-title-main">Euglenales</span> Order of flagellate eukaryotes

Euglenales is an order of flagellates in the phylum Euglenozoa. The family includes the most well-known euglenoid genus, Euglena, as well as other common genera like Phacus and Lepocinclis.

<span class="mw-page-title-main">Euglenaceae</span> Family of flagellate eukaryotes

Euglenaceae is a family of flagellates in the phylum Euglenozoa. The family includes the most well-known euglenoid genus, Euglena.

<span class="mw-page-title-main">Malawimonadidae</span> Family of protists

Malawimonadidae is a family of unicellular eukaryotes of outsize importance in understanding eukaryote phylogeny.

<span class="mw-page-title-main">Jakobid</span>

Jakobids are an order of free-living, heterotrophic, flagellar eukaryotes in the supergroup Excavata. They are small, and can be found in aerobic and anaerobic environments. The order Jakobida, believed to be monophyletic, consists of only twenty species at present, and was classified as a group in 1993. There is ongoing research into the mitochondrial genomes of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of eukaryotes.

Cryomonadida is a group of heterotrophic Rhizaria, that belong to the Cercozoa.

<i>Hemiselmis</i> Genus of single-celled organisms

Hemiselmis is a genus of cryptomonads.

<i>Peranema</i> Genus of protozoans

Peranema is a genus of free-living phagotrophic euglenids. There are more than 20 nominal species, varying in size between 8 and 200 micrometers. Peranema cells are gliding flagellates found in freshwater lakes, ponds and ditches, and are often abundant at the bottom of stagnant pools rich in decaying organic material. Although they belong to the class Euglenoidea, and are morphologically similar to the green Euglena, Peranema have no chloroplasts, and do not conduct autotrophy. Instead, they capture live prey, such as yeast, bacteria and other flagellates, consuming them with the help of a rigid feeding apparatus called a "rod-organ." Unlike the green euglenids, they lack both an eyespot (stigma), and the paraflagellar body (photoreceptor) that is normally coupled with that organelle. However, while Peranema lack a localized photoreceptor, they do possess the light-sensitive protein rhodopsin, and respond to changes in light with a characteristic "curling behaviour."

<i>Neobodo</i> Genus of protists

Neobodo are diverse protists belonging to the eukaryotic supergroup Excavata. They are Kinetoplastids in the subclass Bodonidae. They are small, free-living, heterotrophic flagellates with two flagella of unequal length used to create a propulsive current for feeding. As members of Kinetoplastids, they have an evident kinetoplast There was much confusion and debate within the class Kinetoplastid and subclass Bodonidae regarding the classification of the organism, but finally the new genera Neobodo was proposed by Keith Vickerman. Although they are one of the most common flagellates found in freshwater, they are also able to tolerate saltwater Their ability to alternate between both marine and freshwater environments in many parts of the world give them a “cosmopolitan” character. Due to their relatively microscopic size ranging between 4–12 microns, they are further distinguished as heterotrophic nanoflagellates. This small size ratio limits them as bacterivores that swim around feeding on bacteria attached to surfaces or in aggregates.

Heteronema is a genus of phagotrophic, flagellated euglenoids that are most widely distributed in fresh water environments. This genus consists of two very distinguishable morphogroups that are phylogenetically closely related. These morphogroups are deciphered based on shape, locomotion and other ultrastructural traits. However, this genus does impose taxonomic problems due to the varying historical descriptions of Heteronema species and its similarity to the genus Paranema. The species H. exaratum, was the first heteronemid with a skidding motion to be sequenced, which led to the discovery that it was not closely related to H. scaphrum, contrary to what was previously assumed, but instead to a sister group of primary osmotrophs. This suggests that skidding heteronemids can also be distinguished phylogenetically, being more closely related to Anisoma, Dinema and Aphageae, than to other species within Heteronema.

Postgaardia is a proposed basal clade of flagellate Euglenozoa, following Thomas Cavalier-Smith. As of April 2023, the Interim Register of Marine and Nonmarine Genera treats the group as a subphylum. A 2021 review of Euglenozoa places Cavalier-Smith's proposed members of Postgaardia in the class Symbiontida. As Euglenozoans may be basal eukaryotes, the Postgaardia may be key to studying the evolution of Eukaryotes, including the incorporation of eukaryotic traits such as the incorporation of alphaproteobacterial mitochondrial endosymbionts.

Bihospites is a genus of symbiontid euglenozoans characterized by the presence of two species of epibiotic bacteria on the cell surface. Bihospites cells are clear, biflagellated, and uninucleated, that range between 40–120 μm long and 15–30 μm wide. Bihospites, as well as other members of the symbiontids, are found in semi-anoxic to anoxic sediments in benthic marine environments. Each cell surface is covered by both rod-shaped and spherical-shaped epibiotic bacteria that may share a commensalistic or mutualistic relationship with Bihospites host cells. Bihospites cells are highly contractile and contain several morphological synapomorphies which are present in euglenozoans, however they also contain several unique morphological traits including a unique C-shaped feeding apparatus.

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